A NEW GENUS AND SPECIES OF HERON (AVES: ARDEIDAE) FROM THE LATE MIOCENE OF FLORIDA
David W. Steadman1 and Oona M. Takano2
1Florida Museum of Natural History, University of Florida, Gainesville, Florida 32611-7800 USA <[email protected]>2Department of Biology, University of New Mexico, Albuquerque, NM 87131 USA
Steadman, D. W., and O. M. Takano. 2019. A new genus and species of heron (Aves: Ardeidae) from the late Miocene of Florida. Bulletin of the Florida Museum of Natural History 55(9):174–186.
ABSTRACT
From the recently discovered Montbrook locality, Levy County, Florida (late Miocene; late Hemphillian land mammal age), a complete coracoid and nearly complete scapula represent a large heron that we name Taphophoyx hodgei new genus and species. While the phylogenetic affinities of T. hodgei are not well resolved, the tiger-herons Tigrisoma spp. or boat-billed heron Cochlearius cochlearius (both Neotropical) may be the closest living relative(s) of Taphophoyx, based in large part on several shared characters of the facies articularis clavicularis and facies articularis humeralis. Nevertheless, the coracoid of Taphophoyx has a uniquely prominent facies articularis humeralis and a uniquely sterno-ventral surface of corpus coracoidei. All 21 taxa of birds recorded thus far from Montbrook (mostly aquatic forms such as swans, ducks, geese, grebes, cormorants, ibises, sandpipers, etc.) probably represent extinct species, although Taphophoyx hodgei is the only one assigned to an extinct genus.
Key words: Florida, Montbrook, late Miocene, heron, Ardeidae, new taxa.
This published work and the nomenclatural acts it contains have been registered in ZooBank, the online registration system for the ICZN. The ZooBank Publication number for this issue is BCAF821D-656A-4327-95C1-F2F775F8A959.
Published On-line: April 6, 2019Open Access Download at https://www.floridamuseum.ufl.edu/bulletin/publications/ISSN 2373-9991
Copyright © 2019 by the Florida Museum of Natural History, University of Florida. All rights reserved. Text, images and other media are for nonprofit, educational, or personal use of students, scholars, and the public. Any commercial use or republication by printed or electronic media is strictly prohibited without written permission of the museum.
STEADMAN AND TAKANO: New late Miocene heron from Florida 175
INTRODUCTIONUnderstanding the evolution and historic bioge-ography of North American birds is possible by relating their fossil record to the biochronological system of North American Land Mammal Ages (NALMA), which complements geochronological evidence for the ages of the fossils (Woodburne, 2004). Except for the late Pliocene and Pleisto-cene (Blancan through Rancholabrean NALMAs) occurrences of living species (Lundelius et al., 1983; Webb et al., 2004; Emslie, 1998), the North American fossil record of herons (Ciconiiformes: Ardeidae) is not well developed. From late Mio-cene and early Pliocene (late Clarendonian through late Hemphillian NALMAs) fossil localities, the North American herons described thus far repre-sent extinct species placed in living genera (Becker, 1985; Mayr et al., 2019). Thus, it is of interest that recently discovered late Miocene heron fossils from Florida differ substantially from the compa-rable elements in living genera of herons.
Herons are distributed nearly worldwide today, although they are most diverse in subtropi-cal and tropical regions. A recent compilation rec-ognizes 19 genera and 64 species of herons world-wide (del Hoyo and Collar, 2014), with 8 genera and 12 species currently inhabiting North America. Herons feed primarily in shallow fresh or estuarine waters, where fish, frogs, and crabs are their main source of food.
The heron fossils to be described here are from the Montbrook locality (Florida Museum of Natural History site LV070), south of Williston, Levy County, Florida. Based on occurrences of the rhinoceros Teleoceras, antilocaprid Hexameryx simpsoni, and canid Borophagus hilli, the Mont-brook locality is dated biochronologically to the late Hemphillian (Hh4) NALMA, which would yield an estimated geochronological age of 5.5 to 5 million years ago (Ma; Hulbert, 2018). Among the birds that DWS has identified from Montbrook is the swan Cygnus mariae Bickart, which is oth-erwise known only from the Wickieup locality, Big Sandy Formation, Mohave County, Arizona (Bick-art, 1990). Fission-track (zircon) dating of the Big Sandy Formation yielded ages of 6.1 to 4.6 Ma,
with a mean of 5.5 ± 0.2 Ma (MacFadden et al., 1979), which lends further support to the estimated age of the Montbrook locality.
Discovered in 2015, Montbrook is a sandy alluvial site rather than a sediment-filled fissure/cave/sinkhole as with so many Neogene fossil sites in Florida. In terms of sheer numbers of vertebrate fossils, the Montbrook fauna is dominated by fresh-water fishes, turtles, and alligators. Estuarine fish also are present. The avifauna recovered thus far at Montbrook consists mostly of aquatic birds such as swans, ducks, geese, grebes, cormorants, storks, ibises, and sandpipers (DWS personal observa-tion). Also present are condors, vultures, hawks, galliforms, and passerines. While most of the bird fossils were recovered during excavations, those of smaller taxa have been found through screen-washing of sediments.
MATERIAL AND METHODSAll fossils from Montbrook are catalogued in the Vertebrate Paleontology Collection, Florida Museum of Natural History, University of Florida (UF), Gainesville, Florida, USA. Some fossils from other Florida sites were formerly in the fos-sil collection of Pierce Brodkorb, and now have the prefix UF/PB. Most modern skeletons used in this study are from the UF Ornithology Collec-tion, some of which were formerly in the mod-ern collection of Pierce Brodkorb (PB), as fol-lows. Ardea herodias UF 48756, 50443, 50852, A. cocoi UF 39955, A. cinerea UF 11433, 23037, 23039 (PB 19761), Ardea (Casmerodius) albus UF 42220, 43979, 50858, Egretta rufescens UF 44183, 45453, 51271, E. thula UF 40155, 42736, 43846, 44153, Egretta (Mesophoyx) intermedia UF 23001 (PB 23270), Pilherodius pileatus UF 22906 (PB 26929), Syrigma sibilatrix UF 22940, 38953, Buto-rides virescens UF 40762, 40864, 46769, 48865, Bubulcus ibis UF 40857, 41803, 45498, 45515, Ardeola rufiventris UF 38923, Nyctanassa viola-cea UF 22903 (PB 36755), 42309, 42330, 42737, 49907, Nycticorax nycticorax UF 20838, 22886, 22901 (PB 22328), 40152, Botaurus lentiginosus UF 45502, 48443, 49874, Ixobrychus flavicol-lis UF 39524, I. exilis UF 40336, 43058, 48784,
176 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 55(9)
Tigrisoma mexicanum UF 38926, 42715, T. linea-tum UF 22875 (PB 21581), T. fasciatum UF 43434. We also examined a modern skeleton of Cochle-arius cochlearius from the Museum of Vertebrate Zoology, University of California (MVZ 86641). Measurements were taken using digital calipers with 0.01 mm increments, rounded to the near-est 0.1 mm. The measurements include only one individual for each living species because our goal was merely to give a general idea of the size of the extinct Montbrook heron. Photographs were taken with a Canon EOS 5D Mark II digital camera.
Osteological terminology follows, as closely as possible, that of Baumel and Witmer (1993). We note here that terminology for the coracoid, an essential part of the avian shoulder, can be confus-ing. Four other bones articulate with the coracoid (furcula, humerus, scapula, and sternum, the first three being part of the shoulder). In defining terms, Clark (1993: 12) says “: Regio omalis [Omus] The shoulder is the arbitrarily delimited region of junc-tion of the wing and trunk. The Greek term Omos is used … in its Latin transliteration, Omus.” Among words listed by Brown (1956: 709) for shoulder is “Gr. Omos, m. shoulder, upper arm; akromion, n. point of the shoulder blade; ...” When referring to the end of the coracoid that is opposite from the sternal end, we will call it the “omal” end to sat-isfy reviewers, even though only the furcula articu-lates with the coracoid at this end, separate from where the other two bones of the avian shoulder (humerus, scapula) join the coracoid. In Figures 1–5, we place numbered arrows for each of the 13 diagnostic characters of the new genus and species, except for character 7, which is difficult to discern in photographs.
SYSTEMATIC PALEONTOLOGYOrder PELECANIFORMES1
Family ARDEIDAE LeachTAPHOPHOYX new genus
Type species.—Taphophoyx hodgei new spe-cies.
Zoobank nomenclatural act.—D11FE128-3D90-43A9-BE10-F8C5DFB290F5.1formerly Ciconiiformes.
Diagnosis.—A large heron (Table 1) that differs from all other ardeid genera examined in these characters. CORACOID: 1. In medial aspect, the facies articularis clavicularis is more diagonal (less perpendicular) to long axis of corpus coracoi-dei than in all except Tigrisoma spp. and Cochle-arius cochlearius. 2. In medial aspect, the facies articularis clavicularis lacks a distinct dorsal pro-tuberance (present in all others except Cochlearius cochlearius). 3. In ventral aspect, the sternal end of facies articularis clavicularis is more rounded than in all except Tigrisoma mexicanum, Egretta rufe-scens, and Cochlearius cochlearius. 4. In medio-clavicular aspect, the profile of facies articularis clavicularis is more rounded than in all except Tigrisoma fasciatum, T. lineatum, and Pilherodius pileatus. 5. In lateral, ventral, or dorsal aspect, the sternal margin of facies articularis humeralis is perpendicular to facies articularis scapularis (as in Tigrisoma mexicanum, T. lineatum, and Cochle-arius cochlearius) but joins much more gradually in all others. 6. In ventral aspect, the sternal end of facies articularis humeralis protrudes more abruptly (nearly perpendicularly) from corpus coracoidei. 7. The medio-ventral margin of corpus coracoidei is more rounded (flatter in all others). 8. The depres-sion of sterno-ventral surface of corpus coracoidei is deeper. 9. In medial aspect, the medio-sternal margin of facies articularis sternalis is more con-cave than in all except Casmerodius albus, Egretta rufescens, Mesophoyx intermedia, Nyctanassa violacea, and Cochlearius cochlearius. 10. The medio-ventral surface of facies articularis sternalis is more expanded omally than in all except Botau-rus lentiginosus. SCAPULA: 11. In medial aspect, tuberculum coracoideum blends into facies articu-laris humeralis (separated by distinct notch in all others except Ardea herodias and Tigrisoma mexi-canum). 12. In either medial or lateral aspect, facies articularis humeralis is less protrudent from corpus scapulae than in all except Casmerodius albus and Nyctanassa violacea. 13. In either medial or lateral aspect, facies articularis clavicularis is less pro-trudent proximally than tuberculum coracoideum (more protrudent in all others except Tigrisoma mexicanum and Cochlearius cochlearius).
STEADMAN AND TAKANO: New late Miocene heron from Florida 177
Tabl
e 1.
Mea
sure
men
ts (
in m
m)
of th
e co
raco
id a
nd s
capu
la in
her
ons.
Cc,
cor
pus
cora
coid
ei; f
ac, f
acie
s ar
ticul
aris
cla
vicu
laris
; fah
, fa
cies
arti
cula
ris h
umer
alis
; tc,
tube
rcul
um c
orac
oide
um. D
epth
= d
orso
-ven
tral d
ista
nce;
leng
th =
dis
tanc
e al
ong
ster
no-o
mal
axi
s; w
idth
=
med
io-la
tera
l dis
tanc
e.
Spec
ies
Cor
acoi
dSc
apul
a
Tota
l le
ngth
Dep
th o
f fa
c
Max
le
ngth
of
fah
Max
w
idth
of
fah
Leas
t w
idth
of
cc
Wid
th o
f st
erna
l end
Leng
th
thro
ugh
tc &
fa
h
Wid
th th
roug
h fa
h &
fac
Taph
opho
yx h
odge
i70
.011
.411
.47.
86.
221
.411
.414
.9Ti
gris
oma
mex
ican
um63
.39.
08.
76.
95.
019
.210
.513
.6Ti
gris
oma
linea
tum
58.5
8.1
7.6
5.2
4.4
15.5
8.3
11.0
Tigr
isom
a fa
scia
tum
51.3
7.3
6.4
5.2
4.0
12.8
7.2
10.1
Coc
hlea
rius
coc
hlea
rius
48.0
7.6
7.3
5.0
3.8
14.4
7.8
9.3
Arde
a he
rodi
as74
.713
.812
.38.
06.
224
.112
.316
.0Ar
dea
coco
i72
.211
.911
.67.
76.
120
.611
.916
.7Ar
dea
cine
rea
63.4
11.9
11.3
7.7
5.8
21.6
11.8
14.6
Cas
mer
odiu
s alb
us55
.910
.79.
16.
25.
018
.410
.312
.1Eg
retta
rufe
scen
s47
.68.
58.
35.
94.
117
.28.
69.
9Eg
retta
thul
a36
.96.
57.
04.
43.
412
.86.
38.
3M
esop
hoyx
inte
rmed
ius
41.2
6.6
6.7
4.5
3.6
13.0
7.7
9.8
Pilh
erod
ius p
ileat
us46
.06.
87.
55.
03.
615
.1--
-9.
2Sy
rigm
a si
bila
trix
38.1
7.4
6.2
4.4
3.6
13.7
7.2
8.6
Buto
ride
s vire
scen
s35
.65.
04.
73.
32.
79.
15.
36.
0Bu
bulc
us ib
is37
.66.
16.
53.
83.
111
.16.
97.
7Ar
deol
a ru
fiven
tris
35.1
5.1
4.9
3.4
2.6
9.2
5.0
6.8
Nyc
tana
ssa
viol
acea
48.3
7.8
6.5
4.7
3.9
15.0
7.7
9.0
Nyc
ticor
ax n
yctic
orax
52.7
8.1
7.5
5.6
4.3
15.6
8.4
9.9
Bota
urus
lent
igin
osus
55.1
8.0
6.7
5.7
4.3
15.6
8.4
9.7
Ixob
rych
us fl
avic
ollis
42.8
5.2
4.7
3.6
2.6
10.0
5.1
7.4
178 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 55(9)
Figu
re 1
. The
cor
acoi
d of
her
ons
in d
orsa
l asp
ect.
A. E
gret
ta s
p., U
F/PB
792
5 (f
ossi
l), B
one
Valle
y, F
lorid
a. B
. Ard
ea p
olke
nsis
, UF/
PB
7924
(fos
sil),
Bon
e Va
lley,
Flo
rida.
C. T
apho
phoy
x ho
dgei
new
gen
us a
nd sp
ecie
s, U
F 42
5443
(fos
sil),
Mon
tbro
ok, F
lorid
a. D
. Tig
riso
ma
mex
ican
um, U
F 42
715,
Hon
dura
s. E.
Tig
riso
ma
mex
ican
um, U
F 38
926,
Gua
tem
ala.
F. T
igri
som
a lin
eatu
m, U
F 22
875,
cap
tive.
G. T
igri
-so
ma
fasc
iatu
m, U
F 43
434,
Per
u. H
. Coc
hlea
rius
coc
hlea
rius
, MV
Z 86
641,
El S
alva
dor.
I. Ar
dea
hero
dias
, UF
4875
6, F
lorid
a. J
. Ard
ea
coco
i, U
F 39
955,
Guy
ana.
K. A
rdea
cin
erea
, UF
2303
9, H
olla
nd. L
. Cas
mer
odiu
s al
bus,
UF
4222
0, F
lorid
a. M
. Egr
etta
ruf
esce
ns, U
F 45
453,
Flo
rida.
N. M
esop
hoyx
inte
rmed
ius,
UF
2300
1, Ja
pan.
O. P
ilher
odiu
s pile
atus
, UF
2290
6, c
aptiv
e. P
. Syr
igm
a si
bila
trix
, UF
3895
3,
Arg
entin
a. S
cale
bar
s = 1
0 m
m.
STEADMAN AND TAKANO: New late Miocene heron from Florida 179
Etymology.—Taphophoyx is derived from the Greek words taphos, meaning grave or tomb, and phoyx, meaning heron (Brown, 1956:381, 408). Both words are masculine. The first half of the name Taphophoyx refers to the large concentra-tion of fossils of gomphotheriid proboscideans at Montbrook, rendering the site the paleo-equivalent of an “elephant graveyard.”
Comparisons with other fossil Ardeidae.—Outside of North America, three extinct genera of herons have been described from early Miocene localities (ca. 11–12 million years older than the Montbrook locality); fortunately, each is repre-sented by a partial (omal end) coracoid. The first, Matuku from the St. Bathans Fauna of New Zea-land (~19–16 Ma), was described as not closely related to any particular genus or lineage of heron (Scofield et al., 2010). Matuku differs markedly from Taphophoyx in these characters: 1. In medial aspect, processus acrocoracoideus more pointed (less rounded); 2. In medial aspect, facies articu-laris clavicularis perpendicular to long axis of corpus coracoidei (diagonal in Taphophoyx); 3. In lateral aspect, facies articularis humeralis placed closer to omal end; and 4. In lateral aspect, pro-cessus procoracoideus less protrudent from corpus coracoidei.
The second described early Miocene extinct genus, Pikaihao from the same locality as Matuku, is a small form regarded as allied to the bitterns (Botaurinae; Worthy et al., 2013). Pikaihao differs markedly from Taphophoyx in characters 2 and 4 of Matuku (above), as well as having more pointed (less rounded) omal and sternal margins of facies articularis humeralis.
The third early Miocene genus, Zeltornis from Libya, was a huge heron (larger than in any living species) believed to be related to Cochle-arius or Nycticorax (Balouet, 1981). Known only from the omal half of a coracoid, the measurements of Zeltornis as given by Balouet (1981; translated from the French) are: maximum width of the head – 20.9 mm; minimum width of the head – 12.9 mm; maximum length of the glenoid facet (= facies articularis humeralis) – 17.8 mm. These same three measurements in Taphophoyx are 11.4 mm,
7.3 mm, and 11.6 mm, which range from ~54% to ~65% of the comparable values in Zeltornis. Taphophoyx differs further from Zeltornis in three of the six characters (1, 5, and 6) in the diagnosis of Taphophoyx that can be evaluated.
Other early Miocene heron fossils are of even more uncertain affinities, namely the small Euro-pean forms variously called Proardeola walkeri, Ardea aurelianensis, or Proardea amissa, which represent one or more Ardeola-sized taxon (Olson, 1985; Scofield et al., 2010; Zelenkov, 2011), and Ardeidae sp. from Thailand (Cheneval et al., 1991).
Middle Miocene heron fossils include Ardea sytchevskayae from the Sharga locality in Mongo-lia (~12 Ma), based on a large coracoid (Zelenkov, 2011, 2016), and Ardea sp. from the Sand Canyon Beds (early Barstovian NALMA; ~16–15 Ma) of Nebraska (Becker, 1986), based on a tarsometatar-sus. The coracoid of A. sytchevskayae differs from that of Taphophoyx and is typical of that in Ardea spp. in being slender in dorsal or ventral aspect, in having the facies articularis humeralis located more toward the omal end, and in having the facies articularis clavicularis less diagonal to the long axis of corpus coracoidei. Also from Sharga is a single quadrate described as a new genus, Nycti-soma (Elzanowski and Zelenkov, 2015) with some similarities to Nycticorax, Tigrisoma, and perhaps the extinct Pikaihao. From the Maboko Forma-tion, Kenya (~15–14 Ma), a small ardeid quadrate and tarsometatarsus were referred to cf. Pikaihao (Mayr, 2014; also see Worthy et al., 2013).
Moving to the late Miocene, the new genus and species Ardeagrandis arborea was described from a huge but fragmentary tarsometatarus from Moldova (Kurochkin and Ganya, 1972). All North American late Miocene (late Clarendonian through Hemphillian NALMA) fossils of herons have been referred to living genera, including Ardea, Egretta, Ardeola (Butorides), and Nycticorax (Brodkorb, 1955, 1963; Becker, 1985). We have examined the specimens from localities in Florida (UF 3285, 19001, 21138, 25759, 25939, 25940, 26082), agreeing in each case with Becker’s identifications. Concerning coracoids in particular, the speci-men from Bone Valley, Florida (late Hemphillian,
180 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 55(9)
Figu
re 2
. The
cor
acoi
d of
her
ons i
n m
edia
l asp
ect.
Spec
imen
s as i
n Fi
gure
1. S
cale
bar
s = 1
0 m
m.
STEADMAN AND TAKANO: New late Miocene heron from Florida 181
UF/PB 7924; Figs. 1B, 2B, 3B) referred to Ardea polkensis by Becker (1985) indeed agrees with Ardea rather than Taphophoyx in all seven of the characters (#1–7 above) that can be assessed. Simi-larly, the coracoid from Bone Valley (UF/PB 7925; Figs. 1A, 2A, 3A) that Becker (1985) referred to Egretta sp. agrees with modern Egretta rather than Taphophoyx in all four characters that can be evalu-ated (#4-7 above). The same applies to a previously unreported Bone Valley specimen (UF/PB 9230), which is a similarly sized omal end of a coracoid of Egretta sp.
TAPHOPHOYX HODGEI new speciesHolotype.—Complete left coracoid, UF
425443 (Figs. 1C, 2C, 3C). Montbrook site (LV070), Levy County, Florida. Collected in grid square 15S x 9E by Toni-Ann Benjamin on 9 November 2017.
Paratype.—Left scapula lacking distal part of blade, UF 431654 (Figs. 4A, 5A). Montbrook site (LV070), Levy County, Florida. Collected in grid square 16S x 9E by Sharon Shears on 16 Novem-ber 2017.
Zoobank Nomenclatural Act.—D423D7E9-6D28-49D1-8004-135C342BDAC0.
Species Diagnosis.—As for the genus Tapho-phoyx.
Etymology.—The species name Taphophoyx hodgei is to honor Mr. Eddie Hodge, who has been most generous to the Florida Museum of Natural History in providing logistical help and in granting permission to excavate and study the fossils dis-covered on his land.
Remarks.—The two specimens of Tapho-phoyx hodgei, which are the only heron fossils known from Montbrook, were found in adjacent 1-m2 squares, which suggests that they belong to the same individual. This likelihood increases fur-ther by the fossils being a left coracoid and left scapula, which articulate with each other. Among the 10 characters of the coracoid, Taphophoyx hodgei agrees with Cochlearius cochlearius in five of them, and with one or more species of Tigrisoma in four of them. For the scapula, these agreement values are one of three characters for Cochlearius cochlearius, and two of three characters for Tigri-
soma spp.Compared to those of living species, the mea-
surements of Taphophoyx hodgei are most similar to those of Ardea cinerea and A. cocoi (Table 1). These two large herons have body masses of 1020-2073 g (mean 1443 g) and 1148-1465 g (mean 1306 g), respectively (Dunning, 2008:29).
DISCUSSIONPhylogenetic relationships within the Ardeidae are not well understood, with conflicting proposals based on morphological data, molecular data, and combinations thereof (Payne and Risley, 1976; Sheldon et al., 1995, 2000; McCracken and Sheldon, 1998; Huang et al., 2016; Mayr et al., 2019). We note here that molecular data for phylogenetic analyses are typically based on a single individual tissue sample per species, something that is unavoidable in certain instances, just as with fossils. Thus, we hesitate to place Taphophoyx in a phylogenetic context other than to speculate that, based upon the available coracoid and scapula, it may be more closely related to the tiger-herons (Tigrisoma spp.) or boat-billed heron (Cochlearius cochlearius) than to other living genera of herons. Both of these Neotropical genera occur today as far north as northern Mexico, and likely were more widespread in the Pleistocene (Olson and Suárez, 2008), so the late Neogene existence of a related genus in Florida (and undoubtedly elsewhere in North America) is reasonable.
Thus far, the coracoid and scapula from Montbrook are the only solid evidence of an extinct genus of heron from North America. From the much older (early Miocene; Hemingfordian NALMA; ~18.5 Ma) Thomas Farm locality in Florida, extensive screen-washing of sediments over the past 15 years has produced six fossils (a highly fragmentary coracoid and carpometacarpus, and four pedal phalanges) of a small heron. This material is inadequate for assignment beyond the family level; all other birds from Thomas Farm represented by diagnostic fossils are accommodated in extinct genera (Steadman, 2008).
The late Miocene bird community from Montbrook is dominated by aquatic species such
182 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 55(9)
Figu
re 3
. The
cor
acoi
d of
her
ons i
n ve
ntra
l asp
ect.
Spec
imen
s as i
n Fi
gure
1. S
cale
bar
s = 1
0 m
m.
STEADMAN AND TAKANO: New late Miocene heron from Florida 183
as swans, ducks, grebes, cormorants, herons, and ibises, whether considered from the number of taxa or the number of individual fossils. The Montbrook avifauna does not include the more strictly marine taxa that are so common in the roughly contemporaneous Palmetto Fauna from the Bone Valley phosphate mining district, such as loons, albatrosses, shearwaters, sulids, and alcids (Brobkorb, 1955; Becker, 1987). While many of the birds discovered at Montbrook are compatible with an estuarine environment, the fossils are just as likely to have been deposited in a freshwater setting.
Finally, we note that the Montbrook avifauna
promises to become considerably richer as field and lab work continue. Of the 20+ taxa of birds recognized thus far by DWS, 12 are known from single specimens; we are far from reaching dimin-ishing returns at this unusual site.
ACKNOWLEDGMENTSWe thank the many scientists and volunteers who have excavated and prepared fossils at the Mont-brook site over the past several years. In particu-lar, we thank Toni-Ann Benjamin, Jonathan Bloch, Jason Bourque, Jack Brown, Richard Hulbert, Bill Keeler, Cindy Lockner, Bruce MacFadden, Rachel Narducci, Arthur Poyer, Josh Ringer, Carol and Bill
Figure 4. The scapula of herons in lateral aspect. A. Taphophoyx hodgei new genus and species, UF 431654 (fossil), Montbrook, Florida. B. Tigrisoma mexicanum, UF 42715, Honduras. C. Tigrisoma lineatum, UF 22875, captive. D. Tigrisoma fasciatum, UF 43434, Peru. E. Cochlearius cochlearius, MVZ 86641, El Salvador. F. Ardea herodias, UF 48756, Florida. G. Casmerodius albus, UF 42220, Florida. H. Egretta rufescens, UF 51271, Florida. I. Nyctanassa violacea, UF 22903 (PB 36755), Florida. J. Botaurus lentiginosus, UF 52331, Florida. Scale bars = 10 mm.
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Sewell, Sharon Shears, and Bob Tarnuzzer for their extraordinary efforts. Field work at Montbrook was done with permission and cooperation from Eddie Hodge. Modern skeletal specimens are mainly from UF, supplemented by UCMVZ (through Carla Cicero). This research was supported in part by NSF grants 00127431 and 001756306 (Jona-than Bloch, PI; DWS, co-PI) and by the Felburn Foundation. For comments that improved the man-uscript, we thank two anonymous reviewers, Rich-ard Hulbert, Andrew Kratter, and Arthur Poyer.
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Figure 5. The scapula of herons in medial (costal) aspect. A. Taphophoyx hodgei new genus and species, UF 431654 (fossil), Montbrook, Florida. B. Tigrisoma mexicanum, UF 42715, Honduras. C. Tigrisoma lineatum, UF 22875, captive. D. Tigrisoma fasciatum, UF 43434, Peru. E. Cochlearius cochlearius, MVZ 86641, El Salvador. F. Ardea herodias, UF 48756, Florida. G. Casmerodius albus, UF 42220, Florida. H. Egretta rufescens, UF 51271, Florida. I. Nyctanassa violacea, UF 22903 (PB 36755), Florida. J. Botaurus lentiginosus, UF 52331, Florida. Scale bars = 10 mm.
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